Académique Documents
Professionnel Documents
Culture Documents
CITATIONS READS
218 1,117
2 authors, including:
Vladimir M Zatsiorsky
Pennsylvania State University
350 PUBLICATIONS 10,865 CITATIONS
SEE PROFILE
Some of the authors of this publication are also working on these related
projects:
All content following this page was uploaded by Vladimir M Zatsiorsky on 14 October 2017.
Abstract
Latash, M.L. and V.M. Zatsiorsky, 1993. Joint stiffness: Myth or reality. Human Movement
Science 12. 653-692.
The notion of joint stiffness as commonly studied in biomechanics and motor control is
compared with the physical definition of stiffness. The importance of elastic deformation and
storage of elastic energy is stressed. Different terms are suggested in order to differentiate
between experimentally observed relations between joint angle and torque that are likely to have
different nature. A review of studies measuring stiffness of joint subcomponents and intact joints
is presented. We suggest to either abandon the term ‘joint stiffness’ as misleading or to state up
front stiffness of which of the joint components or subsystems is analyzed in each particular
study. We also suggest that each study of ‘joint stiffness’ should clearly state to what extent the
results are defined by the system’s properties and to what extent they are reflections of the
particular experimental procedure.
1. Introduction
* Corresponding author
section deals with the stiffness of the joint constituent tissues (tendons
and muscles). Since a comprehensive review on the properties of
muscles and tendons has been recently published by Zajac (19891, we
shall just touch these issues and elaborate only on those that have
important implications for control of voluntary movements. In the
third section, joint stiffness is analyzed.
F,= -kx,
k = -AF/Ax, (2)
velocity and directed against the velocity vector, the equation will be:
m d2x/dt2 + b dx/dt + hx = 0, or
d*x/dt* + 1 dx/dt
7
+ WAX= 0, (4)
where m(t) is inertia, b(t) is viscosity, k(t > is stiffness, x is length, and
t is time. A number of researchers have studied the dependencies of
F upon x, and termed the derivative dF/dx stiffness. Let us differen-
tiate both sides of Eq. (5) by t and then divide both sides by dx/dt.
M.L. Latash, V.M. Zatsiorsky / Joint stiffness 657
This equation is quite different from Eq. (2). Its right side is not just
k(t) but a complicated expression. Therefore, using the term ‘stiff-
ness’ for the derivative dF/dx is misleading even if m, b, and k are
not time dependent. This derivative is dependent not only upon time
changes in m, b, and k, but also upon movement kinematics (x, V, A,
and J). That is, it rej7ects both feutures of the system and the method of
testing. In an ideal mechanical system, high inertia and/or viscosity
may lead to high values of d F/dx independently of k(t), and even
when there is no elastic element at all, if the measurements are
performed not at equilibria.
Since dF/dx is frequently used in the literature for describing the
properties of biological objects, let us denote it with a special term
‘quasi-stiffness’, q( t >. By definition:
m
Fig. 1. Two simple physical examples when a derivative dF/dx can be calculated but the noiton
of stiffness is inapplicable. More detail in text.
Let us suggest the following terms for the derivative dF/dx de-
pending upon the physical nature of the system and method of
measurement:
(1) Stiffness (k). The measurements are performed at equilibria. Re-
sistance to the external force is provided by elastic forces, and
potential energy is being stored.
(2) Apparent stiffness (K). The measurements are performed at equi-
libria. The physical nature of the resistive forces is being disre-
garded.
(3) Quasi-stiffness (q). The measurements are performed not at equi-
libria.
Further, we are going to consider systems with lumped parameters
including inertia, viscosity, stiffness, and external forces. That is, in
each case, a simplified mechanical model of the actual system can be
drawn that includes an explicit spring or, more frequently, several
springs. Stiffness will be considered a characteristic of these springs
rather than of the whole system. As we are going to show, the
presence of spring-like components does not assure that the system as
a whole can be attributed a characteristic termed stiffness. If the
models are realistic or too simplistic is another question which is
beyond the scope of the present paper.
It may seem that the main theme of this paper deals mostly with the
problems of terminology rather than with those of motor control or
biomechanics. However, the border is not that strict. First, inappropri-
ate usage of a term borrowed from physics commonly leads to misun-
derstanding if the reader skips the small print in the Methods section.
In extreme cases, it leads to expressions like ‘negative stiffness’
(Dyhre-Poulsen et al., 1991) that do not make physical sense. Second,
a number of contemporary models of motor control, in particular
different versions of the Equilibrium-Point (EP-1 hypothesis, are based
on central regulation of properties of the joints that include joint
stiffness (Feldman, 1980, 1986; Bizzi, 1980; Bizzi et al., 1982, 1992;
660 ML. Lurash, V.M Zutsiortvky / Joint stif@exs
2. I. Tendon stiffness
the tendon fibers rather than their elongation. When a greater tensile
force is applied, the elastic fibers are forcibly stretched, the tendon
stiffness increases and is kept almost constant. The estimates of the
tangential modulus of the stress-strain curve are in the range of
800-1500 MPa for different human and wallaby muscles (Ker et al.,
1986).
In most of the more recent studies, tendon is considered to be
elastic or viscoelastic with a linear elasticity (Ducati et al., 1982;
Oguztoreli and Stein, 1982, 1983). Morgan (1977) has demonstrated
that apparent tendon stiffness is independent of muscle length and
tension induced by electrical stimulation of the motor nerve. Further-
more, Proske and Morgan (1984) have later shown that tendon stiff-
ness remains the same when only parts of the muscle are activated.
On the other hand, changes in tendon stiffness with an increase in
the applied tensile force have been reported by many authors. Rack
and Westbury (1984) stimulated efferent fibers in such a way that
there were virtually no changes in muscle fiber length and all the
movement was supposed to happen in the tendinous component.
Tendon stiffness changed significantly (from 2 to 25 N/mm) with an
increase of muscle force. An increase in tendon stiffness with force
was also shown by Diamant et al. (1972), Ker (19811, Bennett et al.
(19861, and Shadwick (1990).
Note the following mechanical characteristics of tendons:
(a) Energy dissipation in tendons, determined by the area inside the
hysteresis loop curves, is small (Alexander, 1988). It means that
almost all the deformation energy stored in the tendons during
lengthening returns to the system afterwards. Tendons are good
accumulators of the mechanical energy.
(b) The normal physiological range of load lies within the ‘elastic toe’
of the force-deformation curve and is less than 4% of strain which
is considered a limit of tendon reversibility (Harris et al., 1966;
Crisp, 1972).
(cl Above the ‘elastic toe’ range, the tendon stiffness can be regarded
as constant.
clamation marks are added by the authors; compare with Eq. (6) and
the epigraph by Wilkie, 19791. This definition corresponds to quasi-
stiffness as it has been defined earlier. Consequently, caution should
be exercised when data from such muscle fiber studies are used to
describe the mechanical behavior of more complex objects, including
joints.
When a muscle fiber is stimulated, its stiffness (or quasi-stiffness) is
proportional to the overlap between the thick and thin filaments and
changes with time together with fiber tension. In addition, fiber
tension drops to zero when the fiber is permitted to shorten quickly
only 6 nm per half-sarcomere (this distance is short compared to the
size of the crossbridge). Therefore, it has been concluded that in
active muscles, actin and myosin filaments are restricted in their
relative motion and, also, that fiber stiffness results from deformation
within the attached crossbridges. The compliance of the filaments
themselves is very low and, in many cases, may be disregarded (cf.
Jung et al., 1988, 1989).
Stiffness of activated extrafusal muscle fibers increases markedly
(Hoffer and Andreassen, 1981; Proske and Morgan, 1984) to the
extent that it may exceed the stiffness of the serial tendinous compo-
nent (Morgan, 1977; Morgan et al., 1978).
Muscle fiber stiffness is frequently assumed to reside in cross-
bridges and to lead to purely elastic storage of energy. It is assumed to
increase with muscle force (Huxley, 1974; Morgan et al., 1978; Julian
and Sollins, 1975; Walmsley and Proske, 1981; Proske and Morgan,
1984; Rack and Westbury, 1984; Sinkjaer et al., 1988). Based on a
crossbridge model, Morgan (1977) predicted a linear relation between
short-range muscle stiffness and muscle tension. He also experimen-
tally observed such a relation by applying small fast stretches to an
isometrically contracted cat soleus muscle. Short-range stiffness was
found to be dependent upon force but not upon operating length. A
similar conclusion has been reached by Hoffer and Andreassen (1981).
3. Joint stiffness
external load is applied to a body link and the joint angle is measured,
have been used since 1896 (Mosso, 1896; Mosso and Benedicenti,
1896; Rieger, 1906; Reijs, 1921; Filimonoff, 1925; Herzog, 1927;
Spiegel, 1929; Werestchagin, 1931). Vibration measurements were
pioneered by Schalterbrand (1929, 1935, 1937, 1940) and Maikov
(1930) and, then, were used by others (Joyce et al., 1974; Gottlieb and
Agarwal, 1978; MacKay, 1984; MacKay et al., 1986). The underlying
assumption of the method is that the system under consideration can
be described as a linear system with lumped parameters and Eq. (4) is
valid. Passive joint stiffness values, measured in a narrow range of
joint motion, close to the neutral joint position, are usually very low.
They increase sharply near the limits of the joint range of motion.
However, the stretched muscles are typically activated in this range via
the stretch-reflex mechanism, and the joint cannot be considered
passive anymore. Joint flexibility is also influenced by factors different
from stiffness and should be studied as a special phenomenon
(Zatsiorsky, 1977; Hutton, 1992).
Stiffness of relaxed human joints measured from the frequency
response is under 4 Nm/rad for the elbow (Zahalak and Heyman,
1979; Kwan et al., 1979; MacKay et al., 19861, about 1.2 Nm/rad for
the wrist (Lakie et al., 19791, and about 19 Nm/rad for the ankle
(Gottlieb and Agarwal, 1978; Kearney and Hunter, 1982). Note,
however, that in many of these studies, the influence of viscosity upon
natural frequency of the joint was disregarded. Lower values of
passive elbow stiffness of about 1 Nm/rad were reported by Feldman
(1979) based on experiments with smooth unloadings followed by the
joint torque and angle measurements at new equilibria.
Generally, passive joint stiffness:
Let us consider a pin joint with only one degree of freedom. Eq. (3)
will undergo minor transformations:
d2a( t) Wt)
T(t) = m(t)7 + “(t)dt + W)[4t) - %Wl~
where T is torque around the joint, (Y is joint angle, and a<, is resting
angle of the joint. This equation is analyzed with different degrees of
simplifications in most of the mass-spring models of single-joint motor
behavior. Experimental analysis of systems described by Eq. (6) looks
relatively straightforward: An experimenter tries to control time
changes of one of the two measurable variables, torque T(t) or angle
a(t), measures time changes of the other variable, and, with a suffi-
cient number of measurements, is able to calculate time changes of
the coefficients m(t), b(t), and k(t) with some degree of accuracy.
There are numerous ways of performing such experiments including
application of singular small torque perturbations at certain times and
measuring length changes (Ma and Zahalak, 1985; Gottlieb et al.,
1986), using sinusoidal perturbations over a range of frequencies
(Joyce et al., 1974; A garwal and Gottlieb, 1977; Rack et al., 1978;
Cannon and Zahalak, 19821, or using randomized perturbations with
subsequent correlation of torque and angle changes (Hunter and
Kearney, 1982; Lacquaniti et al., 1982; Weiss et al., 1988; Bennett et
al., 1989, 1992).
Attempts at direct application of Eq. (6) for modeling and analysis
of voluntary joint movements lead to considerable complications if
one tries to incorporate at least minimal information about muscles
and their central connections. The first of these problems is that Eq.
(6) models a damped loaded spring with an instantaneous reaction to
external load. Thus, this approach ignores reflex time delays that are
likely to play an important role in movements (especially, fast ones).
Another problem is due to at least two different mechanisms giving
rise to viscous behavior of intact muscles (Feldman et al., 1990). First,
M.L. Latush, KM. Zatsiorsky /Joint stiffness 673
(1) A muscle will oppose the length change by an increase in its force
due to purely mechanical factors, as if it were a rubber band.
(2) There may be a distinct short-latency monosynaptic reaction lead-
ing to a phasic muscle contraction also opposing the external force
but short in duration (Loop 1 in Fig. 3).
(3) There will be a tonic increase in muscle force due to a hypotheti-
cal length-sensitive mechanism commonly associated with the no-
tion of tonic stretch reflex (Loop 2 in Fig. 3).
The lower part of Fig. 3 illustrates all three components as separate
springs with different m, b, and k. Now, imagine that a rapid
‘step-like’ perturbation is applied at a time t, leading to a virtually
instantaneous increase in external torque (load). Muscle length will
start to change. During the first several tens of milliseconds, this
process will depend only on the first factor, i.e. muscle reaction as a
rubber band. After a time delay corresponding to the latency of
monosynaptic reaction for the muscle (t,), there will be a burst of
614 M.L. Latash, KM. Zatsiorsky / Joint stiffness
Force
*
Fig. 3. A schematic drawing of three components that can contribute to spring-like behavior of
an intact muscle (and, consequently, to an intact joint). Inertial and viscous elements are
assumed but not shown. There are time delays in both monosynaptic reflex (Loop 1) and
polysynaptic tonic stretch reflex (Loop 2) arcs.
length changes will reflect properties of only the first of the three
mechanisms providing for spring muscle properties.
It seems absolutely necessary to modify Eq. (8) in order to take into
account the three mechanisms contributing to muscle reaction to
external perturbations. The major problem is not in the presence of
three parallel springs illustrated in Fig. 3, but in their different time
delays. It is also very important to realize what is the cause and what
is the consequence in muscle reactions. If there is an external change
in torque around a joint, it will cause a muscle length change due to
the first, purely peripheral mechanism:
d*a(t) d4)
T(t) - m,,(t)7 -b,,(t)dt = k”(t)bw - a”41 3 (9)
where (Y, corresponds to ‘zero’ angle for the first, peripheral spring.
We will use (Y, and (Ye for ‘zero’ angles for the two other springs. Note
that the viscous component b,(t)(da(t)/d t> in Eq. (9) may be consid-
ered equivalent to Hill’s viscosity (see above).
Changes in muscle length will cause monosynaptic reactions, and
torque changes become a consequence! Taking into account the delay
t,, one gets:
da( t + t2)
-b2(t+t2) dt = k,(t)[4t) - %Wl + W)
tion with the central command which occurs when the afferent signal
comes to the hypothetical central processing unit. This factor can
easily be introduced by changing the argument of cu,(t> and k,(t)
functions:
da( t + t2)
-b2(t+t2) dt = W)b(t) - a,(t)1 + W)
The methods for analysis of Eq. (14) have certainly been based on a
number of assumptions and simplifications. It is important, however,
to make sure that these simplifications are explicitly stated, their
possible effects on the quality of the results are assessed, and they do
not eliminate the basic features of the hypothetical motor control
process.
(1) Muscles have central reflex connections that provide for delayed
spring-like reactions.
(2) Stiffness and viscosity in each of the hypothetical loops are likely
to be length-, velocity-, and time-dependent.
5. Concluding remarks
’ Recently, MacMahon (1990) suggested to differentiate two stiffnesses during the support
phase in locomotion, one due to knee bending, and the other due to leg rotation. Leg rotation
can be compared to an inverted pendulum rotation. Therefore, there will be changes of the ratio
of vertical component of ground reaction force to vertical displacement of the body center of
gravity (cf. Fig. l).However, the notion of stiffness can hardly be applied to such phenomena.
M.Ld. Latash, VM. Zatsiorslcy /Joint stiffness 683
bodies are supposed to maintain constant shape. Muscles are not such
bodies, and joints are complex system consisting of several bodies.
(2) The following terms for the derivative dF/dx are suggested:
(a) Stiffness (k). The measurements are performed at equilibria.
Resistance to the external force is provided by elastic forces, and
potential energy is being stored.
(b) Apparent stiffness (K). The measurements are performed at
equilibria. The physical nature of the resistive forces is being disre-
garded.
(c) Quasi-stiffness (4). The measurements are performed not at
equilibria.
Apparent stiffness (K) and quasi-stiffness (4) are easier to measure.
However, these indices do not reflect participation of elastic forces
and accumulation of elastic energy. In particular, 9 can easily be
negative while genuine stiffness is always positive. We suggest that
studies involving experimental procedures that lead to calculation of K
or 4 should clearly state the difference between these measures and
the physical notion of stiffness.
(3) Stiffness measurement of passive biological tissues (tendons,
muscles, and passive joints), albeit not easy, does not meet with
conceptual difficulties.
(4) In studies of active muscle fibers, a wealth of important scien-
tific knowledge has been collected with methods of small stretches
and releases. However, these methods measure quasi-stiffness (or, at
best, apparent stiffness) of muscle fibers. Consequently, caution should
be exercised when data from these studies are used to describe the
mechanical behavior of more complex objects, joints included.
(5) Muscle stiffness is determined, according to the well-known Hill
model, by its parallel (PEC) as well as series (SEC) elastic compo-
nents. Vibration methods for measuring muscle stiffness in humans
are based on several assumptions which are in contradiction with the
Hill’s model. Consequently, ‘stiffness’ measurements based on two
different models provide different results.
(6) The muscle-tendon complex cannot be considered a single
spring and, therefore, cannot be assigned a value of stiffness. In
general, elastic energy storage in muscle-tendon complexes resides
primarily in tendons rather than in muscles. However, depending on
the type of the muscle, level of muscle activation, operating length of
the muscle-tendon complex, and parameters of external perturbation,
resistance to the perturbation will receive different contributions from
the tendon serial compliance and compliance of muscle fibers.
(7) Passive joint stiffness is determined when all muscles crossing
the joint are relaxed. Passive joint stiffness: (a) is higher at a small
range of limb displacement (short-range stiffness); (b) depends on the
initial joint position; cc> decreases over a period of 1 s due to viscous
relaxation (although, this comment refers to frequently measured
quasi-stiffness); (d) depends on the direction of joint motion.
(8) An intact joint is a complex, non-linear system whose angle
changes under a changing external force cannot be described with just
one parameter. One can probably take into account the inertial
component with a reasonable degree of accuracy. The presence of two
components in the velocity-dependent force generation by the muscle
(see above), one of which has a reflex time delay, makes analysis of
viscous forces quite complicated. The same is true for the elastic
components of muscle reaction. Therefore, we feel that experimental
analysis is very likely to lead to assessments of joint stiffness and
viscosity dependent upon both properties of the studied system and
particular method used for making these assessments. Therefore, we
think that intact human joint cannot be assigned a parameter termed
‘stiffness’.
(9) This does not mean, however, that stiffness (and viscosity) of
joint subsystems and subcomponents cannot be studied. Considerable
success in the studies of passive structures suggests that at least some
of the joint subcomponents can be assigned values of stiffness (and
viscosity) that comply with the strict definitions introduced in physics.
We also feel optimistic towards the introduced method of study of
stiffness of the tonic stretch reflex subcomponent that contributes to
the overall joint behavior and is assumed to play a major role in the
process of single-joint motor control. We believe that all the other
subcomponents should be studied separately from each other by
designing special experimental manipulations that are able to separate
them according, for example, to their characteristic time constants.
(10) We suggest to either abandon the term ‘joint stiffness’ as
misleading or to state upfront stiffness of which of the joint compo-
nents or subsystems is analyzed in each particular study. We also
suggest that each study of ‘joint stiffness’ should clearly state to what
extent the results are defined by the system’s properties and to what
extent they are reflections of the particular experimental procedure.
M.L. Latash, KM. Zatsiorsky / Joint stiffness 685
Acknowledgments
The authors express their gratitude to Dr. Ziaul Hasan for his
profound and helpful comments. Preparation of this manuscript was
in part supported by an NIH grant HD 30128.
References
Abdusamatov, R.M. and A.G. Feldman, 1986. Description of the electromyograms with the aid
of a mathematical model for single joint movements. Biophysics 31, 549-552.
Agatwal, G.C. and G.L. Gottlieb, 1977. Compliance of the human ankle joint. Journal of
Biomechal Engineering 99, 166-170.
Alexander, R.McN., 1981. ‘Mechanics of skeleton and tendons’. In: B.B. Brooks (ed.), Handbook
of physiology. Section a. The nervous system. Vol. 2 Motor control (pp. 17-42). Bethesda,
MD: American Physiological Society.
Alexander, R.McN., 1988. Elastic mechanisms in animal movement. Cambridge: Cambridge
University Press.
Alexander, R.McN. and H.C. BennettClark, 1977. Storage of elastic strain energy in muscle and
other tissue. Nature 265, 114-l 17.
Alexander, R.McN. and A. Vernon, 1975. Mechanics of hopping by kangaroos (Macropodidae).
Journal of Zoology (London) 177, 265-303.
Alexander, R.McN., G.M.O. Maloiy, R.F. Ker, AS. Jayes and C.N. Warui, 1982. The role of
tendon elasticity in the locomotion of the camel (Camelus dromedarius). Journal of Zoology
(London) 198, 2933313.
Alnaqeeb, M.A., N.S. Al Zaid and G. Goldspink, 1984. Connective tissue changes and physical
properties of developing and ageing skeletal muscle. Journal of Anatomy 139, 6777689.
Arndt, K.H., 1976. Achillesehnenruptur and Sport. Leipzig: Johann Ambrosius Barth.
Aruin. A.S., V.M. Zatsiorsky, L.M. Raitsin, N.I. Volkov and E.A. Shirkovets, 1977. Influence of
elastic forces of the muscles upon the efficiency of muscle work. Human Physiology 3,
420-426.
Aruin, AS., V.M. Zatsiorsky, G.J. Panovko and L.M. Raitsin, 1979. Equivalent biomechanical
characteristics of the ankle joint muscles. Human Physiology 4, 862-868.
Asatryan, D.G. and A.G. Feldman, 1965. Functional tuning of the nervous system with control of
movements or maintenance of a steady posture. I. Mechanographic analysis of the work of
the limb on execution of a postural task. Biophysics 10, 925-935.
Bahler, A.S., 1967. Series elastic component of mammalian skeletal muscle. American Journal of
Physiology 213, 1560-1564.
Bennett, M.B., R.F. Ker, N.J. Dimery and R.McN. Alexander, 1986. Mechanical properties of
various mammalian tendons. Journal of Zoology (London) 209, 5377548.
686 M.L. Latash, VM. Zatsiorsky / Joint stiffness
Bennett, D.J., Y. Xu, J.M. Hollerbach and I.W. Hunter, 1989. Identifying the mechanical
impedance of the elbow joint during posture and movement. Abstracts of the Society for
Neuroscience 15, 396.
Bennett, D.J., J.M. Hollerbach, Y. Xu and I.W. Hunter. 1992. Time-varying stiffness of human
elbow joint during cyclic voluntary movement. Experimental Brain Research 88, 433-442.
Berkwitz, NJ., 1932. Quantitative studies on the human muscle tonus. II. An analysis of
eighty-two normal and pathological cases. Archives of Neurology and Psychiatry 28, 603-614.
Bernstein, N.A., 1947. On the construction of movements. Moscow: Medgiz (in Russian).
Bernstein, N.A., 1967. The co-ordination and regulation of movements. Oxford: Pergamon Press.
Bigland, B. and 0. Lippold, 1954. The relation between force, velocity and integrated electrical
activity in human muscles. Journal of Physiology 123, 214-224.
Bizzi, E., 1980. ‘Central and peripheral mechanisms in motor control’. In: G.E. Stelmach and J.
Requin (eds.), Tutorials in motor behavior (pp. 131-143). Amsterdam: North-Holland.
Bizzi, E., N. Accornero, W. Chapple and N. Hogan, 1982. Arm trajectory formation in monkeys.
Experimental Brain Research 46, 139-143.
Bizzi, E., N. Hogan, F.A. Mussa-Ivaldi and S. Giszter. 1992. Does the nervous system use
equilibrium-point control to guide single and multiple joint movements? Behavioral and
Brain Sciences 15, 603-613.
Blanpied, P. and G.L. Smidt, 1992. Human plantarflexor stiffness to multiple single-stretch trials.
Journal of Biomechanics 25, 29-39.
Bobbert, M.F.. P.A. Huijing and G.J. van Ingen Schenau, 1986. A model of the human triceps
surae muscle-tendon complex applied to jumping. Journal of Biomechanics 19, 887-898.
Borg, T.K. and J.B. Caulfield, 1980. Morphology of connective tissue in skeletal muscle. Tissue
and Cell 12, 197-207.
Brenner, B., 1990. ‘Muscle mechanics and biochemical kinetics’. In: J.M. Squire (ed.), Molecular
mechanics in muscular contraction (pp. 77-150). Boca Raton, FL: CRC Press.
Broman, T., 1949. Electra-mechanographic registrations of passive movements in normal and
pathological subjects. Acta Psychiatrica et Neurologica Suppl. 53, 1.
Buchanan, T.S., D.P.J. Almdale. J.L. Lewis and W.Z. Rymer. 1986. Characteristics of synergetic
relations during isometric contractions of human elbow muscles. Journal of Neurophysiology
56, 1225-1241.
Buchtal, F., 1957. Introduction to electromyography. Kobenhavn: Stand. University.
Buchtal, F. and S. Clemmesen, 1946. Action potentials in pathological postural reflex activity
(spacticity and rigidity). Acta Psychiatrica et Neurologica 21, 151-162.
Cannon, S.C. and G.I. Zahalak, 1982. The mechanical behavior of active human skeletal muscle
in small oscillations. Journal of Biomechanics 15, 111-121.
Cordo, P.J. and W.Z. Rymer, 1982. Motor unit activation patterns in lengthening and isometric
contractions of hindlimb extensor muscles in the decerebrate cat. Journal of Neurophysiology
47, 782-796.
Crago, P.E., J.C. Houk and Z. Hasan, 1976. Regulatory actions of human stretch reflex. Journal
of Neurophysiology 39, 925-935.
Crisp, J.D.C., 1972. ‘Properties of tendon and skin’. In: Y.C. Fung. N. Perrone and M. Anliker
(eds.). Biomechanics. Its foundations and objectives (pp. 141&180). Englewood Cliffs, NJ:
Prentice-Hall.
Davis, W.R. and J.A.S. Kelso, 1982. Analysis of ‘invariant characteristics’ in the motor control of
Down’s syndrome and normal subjects. Journal of Motor Behavior 14, 194-212.
De Serre, S.J. and T.E. Milner. 1991. Wrist muscle activation patterns and stiffness associated
with stable and unstable mechanical loads. Experimental Brain Research 86. 451-458.
Diamant, J., A. Keller, E. Baer, M. Litt and R.G.C. Arridge, 1972. Collagen: Ultrastructurre and
its relation to mechanical properties as a function of ageing. Proceedings of the Royal Society
B 180, 239-315.
M.L. Latash, V.M. Zatsiorsky / Joint stiffness 687
Ducati, A., F. Parmiggiani and M. Scieppati, 1982. Simulation of post-tetanic potentiation and
fatigue in muscle using a visco-elastic model. Biological Cybernetics 44, 129-135.
Dyhre-Paulsen, P., E.B. Simonsen and M. Voigt, 1991. Dynamic control of muscle stiffness and
H reflex modulation during hopping and jumping in man. Journal of Physiology 437,
287-304.
Elliot, D.H., 1965. Structure and function of mammalian tendon. Biological Reviews 40,
3922421.
Encyclopaedia of Science and Technology, 1977. Impedance, Mechanical, 7 (p. 44). New York:
McGraw-Hill.
Feldman, A.G., 1966. Functional tuning of the nervous system with control of movement or
maintenance of a steady posture. II. Controllable parameters of the muscle. Biophysics 11,
565-578.
Feldman, A.G., 1979. Central and reflex mechanisms of motor control. Moscow: Nauka (in
Russian).
Feldman, A.G., 1980. Superposition of motor programs. I. Rhythmic forearm movements in
man. Neuroscience 5, 81-90.
Feldman, A.G., 1986. Once more on the equilibrium-point hypothesis (I model) for motor
control. Journal of Motor Behavior 18, 17-54.
Feldman, A.G., S.V. Adamovitch, D.J. Ostty and J.R. Flanagan, 1990. ‘The origin of electromyo-
grams - Explanations based on the equilibrium point hypothesis’. In: J.M. Winters and
S.L.-Y. Woo teds.), Multiple muscle systems. Biomecuanics and movement organization (pp.
195-213). Berlin: Springer-Verlag.
Feldman, A.G. and M.L. Latash, 1982. Afferent and efferent components of joint position sense:
Interpretation of kinaesthetic illusions. Biological Cybernetics 42, 2055214.
Fenn, W.O. and P.H. Carvey, 1934. The measurement of the elasticity and viscosity of skeletal
muscle in normal and pathological cases; a study of so-called ‘muscle tonus’. Journal of
Clinical Investigations 19, 383-397.
Filimonoff, J.N., 1925. Klinische Beitrage zum Tonusproblem. Zeitschrift fur die gesamte
Neurologie und Psychiatric 96, 368-384.
Gleim, G., N. Stachefeld and J. Nicholas, 1990. The influence of flexibility on the economy of
walking and jogging. Journal of Orthopaedic Research 8, 814-823.
Gottlieb, G.L. and G.C. Agarwal, 1978. Dependence of human ankle compliance on joint angle.
Journal of Biomechanics 11, 177-181.
Gottlieb, G.L. and G.C. Agarwal, 1988. Compliance of single joints: Elastic and plastic charac-
teristics. Journal of Neurophysiology 59, 937-951.
Gottlieb, G.L., G.C. Agarwal and A.S. Tanavde, 1986. A method of estimating human joint
compliance which is insensitive to reflex, triggered or voluntary reactions. Abstracts of the
Society for Neuroscience 12, 468.
Goubel, F., 1974. Les proprietes mechaniques du muscle au tours du mouvement sous-maximal
(p. 217). These Doctorat d’Etat, Fat. Sci. Lille, France.
Goubel, F. and E. Pertuzon, 1973. Evaluation de I’elasticite du muscle in situ par une methode
de quick-release. Archives Internationales de Physiologie et Biochimie 81, 697-701.
Greene, P.R. and T.A. MacMahon, 1979. Reflex stiffness of man’s antigravity muscles during
kneebends while carrying extra weights. Journal of Biomechanics 12, 881-891.
Griffiths, R.I., 1984. Mechanical properties of an ankle extensor muscle in a freely hopping
wallaby. Ph.D. Thesis. Monash Univ. Victoria, Australia.
Griffiths, R.I., 1987. Ultrasound transit time gives direct measurement of muscle fibre length in
vivo. Journal of Neuroscience Methods 21, 159-165.
Griffiths, R.I., 1991. Shortening of muscle fibres during stretch of the active cat medial
gastrocnemius muscle: The role of tendon compliance. Journal of Physiology 436, 219-236.
688 M.L. Latash, V.M. Zatsiorsky / Joint stiffness
Gurfinkel, VS., Y.M. Kots and M.L. Schick, 1965. Control of body posture. Nauka, Moscow (in
Russian).
Hasan, Z., 1992. Is stiffness the mainspring of posture and movement? Behavioral and Brain
Sciences 15, 7566758.
Hasan, Z. and R.M. Enoka, 1985. Isometric torque-angle relationship and movement-related
activity of human elbow flexors: Implications for the equilibrium-point hypothesis. Experi-
mental Brain Research 59, 441-450.
Heerkens, Y.F., R.D. Woittiez, P.A. Huijing, A. Huson and G.J. van Ingen Schenau, 1981.
Inter-individual differences in passive resistance of the human knee. Human Movement
Science 4, 167-188.
Herzog, F., 1927. Uber die myotatische Innervation antagonistischer Muskeln. Deutsche
Zeitschrift fin die Nervenheilkunde 96, 22-37.
Hill. A.V., 1938. The heat of shortening and the dynamic constants of muscle. Proceedings of the
Royal Society of London B 126, 1366195.
Hill, A.V., 1949. The heat of activation and the heat of shortening in a muscle twitch.
Proceedings of the Royal Society of London B 136, 195-211.
Hill, A.V., 1950. The series elastic components of muscle. Proceedings of the Royal Society of
London B 137, 2733280.
Hill, A.V., 1953. The mechanics of active muscle. Proceedings of the Royal Society of London B
141, 104-117.
Hill, A.V., 1960. Production and absorbtion of work by muscle. Science 131, 897-903.
Hill, A.V.. 1970. First and last experiments in muscle mechanics. Cambridge: Cambridge
University Press.
Hoefer, P.F.A. and J.J. Putnam, 1939. Action potentials of muscles in normal subjects. Archives
of Neurology and Psychiatry 42, 201-210.
Hof, A.L., 1990. ‘Effects of muscle elasticity in walking and running’. In: J.M. Winters and
S.L.-Y. Woo teds.), Multiple muscle systems: Biomechanics and movement organization (pp.
591-607). Berlin: Springer-Verlag.
Hof. A.L. and J.W. van den Berg, 1981. EMG to force processing. II. Estimation of parameters
of the Hill muscle model for the human triceps surae by means of a calf ergometer. Journal
of Biomechanics 14. 759-770.
Hof, A.L.. B.A. Geelan and J. van den Berg, 1983. Calf muscle movement. work and efficiency in
level walking; roles of series elasticity. Journal of Biomechanics 16, 5233537.
Hoffer, J.A. and S. Andreassen, 1981. Regulation of soleus muscle stiffness in premammillary
cats: Intrinsic and reflex components. Journal of Neurophysiology 45, 267-285.
Hoffer, J.A., A.A. Caputi, I.E. Pose and RI. Griffiths, 1989. Roles of muscle activity and load on
the relationship between muscle spindle length and whole muscle length in the freely walking
cat. Progress in Brain Research 80, 75085.
Hogan, N., 1990. ‘Mechanical impedance of single- and multi-articular systems’. In: J.M. Winters
and S.L.-Y. Woo teds.), Multiple muscle systems: Biomechanics and movement organization
(pp. 149-164). Berlin: Springer-Verlag.
Huijing, P.A., 1992. ‘Elastic potential of muscle’. In: P.V. Komi ted.), Strength and power in
sport (pp. 151-168). Oxford: Blackwell.
Hunter, I.W. and R.E. Kearney, 1982. Dynamics of human ankle stiffness: Variation with mean
ankle torque. Journal of Biomechanics 15, 747-752.
Hutton, R.S., 1992. ‘Neuromuscular basis of stretching exercises’. In: P.V. Komi (ed.1, Strength
and power in sport. The encyclopaedia of sports medicine (pp. 29-38). Oxford: Blackwell.
Huxley, A.F., 1974. Review lecture: muscular contraction. Journal of Physiology 243, l-43.
Huxley, A.F. and R.M. Simmons, 1970. Rapid ‘give’ and the tension ‘shoulder’ in the relaxation
of frog muscle fibres. Journal of Physiology 210, 32233.
M.L. L&ash, V.M. Zatsiorsky / Joint stiffness 689
Jewell, B.R. and D.R. Wilkie, 1958. An analysis of the mechanical components of frog’s striated
muscle. Journal of Physiology 143, 515-540.
Joyce, G.C., P.M.H. Rack and D.R. Westbury, 1969. The mechanical properties of cat soleus
muscle during controlled lengthening and shortening movements Journal of Physiology 204,
461-474.
Joyce, G.C., P.M.H. Rack and H.F. Ross, 1974. The forces generated in the human elbow joint
in response to imposed sinusoidal movements of the forearm. Journal of Physiology 240,
351-374.
Julian, F.J. and M.R. Sollins, 1975. Variation of muscle stiffness with force at increasing speeds
of shortening. Journal of General Physiology 66, 287-302.
Jung, D.W.G., T. Blangi, H. de Graaf and B.W. Treijtel, 1988. Elastic properties of relaxed,
activated, and rigor muscle fibers measured with microsecond resolution. Biophysical Journal
54, 897-908.
Jung, D.W.G., T. Blange, H. de Graaf, and B.W. Treijtel, 1989. Weakly attached cross-bridges in
relaxed frog muscle fibers. Biophysical Journal 55, 6055619.
Kearney, R.E. and I.W. Hunter, 1982. Dynamics of human ankle stiffness: Variation with
displacement amplitude. Journal of Biomechanics 15, 753-756.
Ker, R.F., 1981. Dynamic tensile properties of the plantaris tendon of sheep (Ovis aries). Journal
of Experimental Biology 93, 283-302.
Ker, R.F., N.J. Dimery and R.McN. Alexander, 1986. The role of tendon elasticity in hopping in
a wallaby (Macropus rufogriseus). Journal of Zoology (London) 208, 417-428.
Komi, P.V., 1992. ‘Stretch-shortening cycle’. In: P.V. Komi (ed.), Strength and power in sport
(pp. 169-179). Oxford: Blackwell.
Kulig, K., J. Andrews and J.G. Hay, 1984. Human strength curves. Exercise and Sport Science
Reviews 12, 417-466.
Kwan, H.C., J.T. Murphy and M.W. Repeck, 1979. Control of stiffness by the medium latency
electromyographic response to limb perturbation. Canadian Journal of Physiology and
Pharmacology 57, 277-285.
Lacquaniti, F., F. Licata and J.F. Soechting, 1982. The mechanical behavior of the human
forearm in response to transient perturbations. Biological Cybernetics 44, 67-77.
Lakie, M., E.G. Walsh and G. Wright, 1979. Wrist compliance. Journal of Physiology 295,
9x-99P.
Lakie, M., E.G. Walsh and G. Wright, 1981. Measurements of inertia of the hand, and the
stiffness of the forearm muscles using resonant frequency methods, with added inertia or
position feedback. Journal of Physiology 310, 3-4P.
Lan, N. and P.E. Crago, 1992. Equilibrium-point hypothesis, minimum effort control strategy
and the triphasic muscle activation pattern. Behavioral and Brain Science 15, 769-771.
Latash, M.L., 1992. Virtual trajectories, joint stiffness, and changes in natural frequency during
single-joint oscillatory movements. Neuroscience 49, 209-220.
Latash, M.L. and G.L. Gottlieb, 1990. Compliant characteristics of single joints: Preservation of
equifinality with phasic reactions. Biological Cybernetics 62, 331-336.
Latash, M.L. and G.L. Gottlieb, 1991a. Reconstruction of joint compliant characteristics during
fast and slow movements. Neuroscience 43, 697-712.
Latash, M.L. and G.L. Gottlieb, 1991b. An equilibrium-point model of dynamic regulation for
fast single-joint movements: 1. Emergence of strategy-dependent EMG patterns. Journal of
Motor Behavior 23, 163-177.
Lehman, B., 1940. Zur Physiologie des Liegens. Arheitsphysiologie 11, 253-264.
Lestienne, F. and E. Pertuzon, 1974. Determination, in situ, de la visco-elasticit& du muscle
humain inactive. European Journal of Applied Physiology 32, 159-170.
Ma, S.-P. and G.I. Zahalak, 1985. The mechanical response of the active human triceps brachii
muscle to very rapid stretch and shortening. Journal of Biomechanics 18, 585-598.
690 M.L. Latash, KM. Zutsiorsky / Joint stiffness
MacKay, W.A., 1984. Resonance properties of the human elbow. Canadian Journal of Physiol-
ogy and Pharmacology 62, 802-808.
MacKay, W.A., D.J. Crammond, H.C. Kwan and J.T. Murphy, 1986. Measurements of human
forearm viscoelasticity. Journal of Biomechanics 19, 231-238.
McKinley, J.C. and N.I. Berkwitz, 1928. Quantitative studies on human muscle tonus. Archives
of Neurology and Psychiatry 19, 1036-1045.
McKinley, J.C. and N.I. Berkwitz. 1933. Electric action potentials in muscles during recording of
mechanical tonus tracing. Archives of Neurology and Psychiatry 29, 272-280.
MacMahon, T.A., 1984. Muscles. reflexes, and locomotion. Princeton, NJ: Princeton University
Press.
MacMahon, T.A., 1990. ‘Spring-like properties of muscles and reflexes in running’. In: J.M.
Winters and S.L.-Y. Woo (eds.), Multiple muscle systems. Biomechanics and movement
organization (pp. 380-390). Berlin: Springer-Verlag.
MacMahon, T.A. and P.R. Greene, 1979. The influence of track compliance in running. Journal
of Biomechanics 12, 893-904.
McPherson, L., 19.53. A method of determining the force-velocity relation of muscle from two
isometric contractions. Journal of Physiology 122, 172-177.
Magid, A. and D.J. Law, 1985. Myofibrils bear most of the resting tension in frog skeletal
muscle. Science 230, 1280-1282.
Maikov, S.S., 1930. Study of muscular tonus. Reviews in Psychiatry, Neurology and Reflexology
3-4, 44-55 (in Russian).
Morgan, D.L., 1977. Separation of active and passive components of short-range stiffness of
muscle. America1 Journal of Physiology 232, C45-C49.
Morgan, D.L., U. Proske and D. Warren. 1978. Measurements of muscle stiffness and the
mechanism of elastic storage of energy in hopping kangaroos. Journal of Physiology 282.
253-261.
Mosso, A., 1896. Description d’un myotonometre pour Ctudier la tonicite des muscles chez
I’homme. Italian Biological Archive 25, 385.
Mosso, A. and A. Benedicenti. 1896. La tonicite des muscles Ctudiee chez I’homme. Italian
Biological Archive 25, 385.
Oguztoreli, M.N. and R.B. Stein, 1982. Analysis of a model for antagonistic muscles. Biological
Cybernetics 45, 177-184.
Oguztoreli, M.N. and R.B. Stein, 1983. Optimal control of antagonistic muscles. Biological
Cybernetics 48, 91-99.
Pertuzon, E., 1968. Un dispositif pour les experiences de quick release. Le Travail Humaine 31,
303-308.
Pertuzon, E., 1972. La contraction musculaire dans le mouvement volontaire maximal (p. 208).
These Doctorat d’Etat, Fat. Sci., Lille, France.
Piziale. R.L. and J.C. Rastegar, 1977. Measurement of the non-linear, coupled stiffness charac-
teristics of the human knee. Journal of Biomechanics 10, 45-51.
Proske, U. and D.L. Morgan, 1984. Stiffness of cat soleus muscle and tendon during activation of
part of muscle. Journal of Neurophysiology 52, 459-468.
Proske, U. and D.L. Morgan, 1987. Tendon stiffness: Methods of measurement and significance
for the control of movement. A review. Journal of Biomechanics 20, 75-82.
Rack, P.M.H., 1981. ‘Limitations of somatosensory feedback in control of posture and move-
ment’. In: V.B. Brooks (ed.), Handbook of physiology, section 1: The nervous system, vol. 2:
Motor control (pp. 229-256). New York: American Physiological Society.
Rack, P.M.H. and H.F. Ross, 1984. The tendon of flexor pollicis longus: Its effects on the
muscular control of force and position at the human thumb. Journal of Physiology 351,
999110.
M.L. Latash, KM. Zatsiorsky / Joint stiffness 691
Rack, P.M.H. and D.R. Westbury, 1984. Elastic properties of the cat soleus tendon and their
functional importance. Journal of Physiology 347, 479-495.
Rack, P.M.H., H.F. Ross and T.I.H. Brown, 1978. ‘Reflex responses during sinusoidal movement
of human limbs’. In: J.E. Desmedt ted.), Cerebral Motor control in man: Long loop
mechanisms (pp. 216-118). Base]: Karger.
Rack. P.M.H., H.F. Ross, A.F. Thilmann and D.K.W. Walters, 1983. Reflex responses at the
human ankle: The importance of tendon compliance. Journal of Physiology 347, 479-495.
Ralston, H.J., V.T. Inman, L.A. Strait and M.D. Shaffrath, 1947. Mechanics of human isolated
voluntary muscle. American Journal of Physiology 151, 612-620.
Ralston, H.J. and B. Libet, 1953. The question of tonus in skeletal muscle. American Journal of
Physical Medicine 2, 85-91.
Reichel, H., A. Bleichert and F. Zimmer, 1956. Die elastischen Eigenschaften des Skelets and
Herzmuskels. Zeitschrift fir Biologie 108, 188-195.
Reijs, I.H.O., 1921. Uber die Veranderung der Kraft wahrend der Bewegung. Pfliiger’s Archiv
fir die gesamte Physiologie 91, 2344249.
Rieger, C., 1906. Untersuchungen iiber Muskelzustande. Jena
Ritchie, J.M. and D.R. Wilkie, 1958. The dynamics of muscular contraction. Journal of
Physiology 143, 104-124.
Sandow, A., 1954. On the active state of striated muscle. Science 127, 760-769.
Schalterbrand, G., 1929. Messung des Dehnungswiderstandes am menschlichen Muskel bei
Gesunden, Spastikern und Parkinsonismusfallen. Deutsche Zeitschrift fir die Nerven-
heilkunde 109, 231-250.
Schalterbrand, G., 1935. Verhalten der myotatischen Reflexe bei verschiedenen Tonusstorungen.
Deutsche Zeitschrift fir die Nervenheilkunde 136, l-18.
Schalterbrand, Cl., 1937. Myographische Untersuchungen in der Klinik. 1. Geschichte der
Myographie und Beschreibung eines neuen Myographen. Deutsche Zeitschrift fur die Ner-
venheilkunde 142, l-24.
Schalterbrand, G., 1940. Die Muskelspannungsmessung (Myographie) und ihre Bedeutung far
die klinische Diagnostik. Zentralblatt fir die innere Medizin 61, 1-25.
Shabashova, A.S., 1947. Balance of tonus of antagonistic muscle groups. PH.D. thesis, Moscow
(Cited after Gurfinkel, Kots and Shick, 1965).
Shadwick, R.E., 1990. Elastic energy storage in tendons: Mechanical differences related to
function and age. Journal of Applied Physiology 68, 1033-1040.
Sherrington, ChS., 1906. Integrated activity of the nervous system. New Haven, CT: Yalen
University Press.
Sinkjaer, T., E. Toft, S. Andreassen and B.C. Hornemann. 1988. Muscle stiffness in human ankle
dorsiflexors: Intrinsic and reflex components. Journal of Neurophysiology 60, 11 IO- 1121.
Spiegel, E.A., 1929. Zur Methodik der Tonusuntersuchung am Menschen. Zeitschrift fir die
gesamte Neurologie und Psychiatric 122, 475-484.
Such, C.H., A. Unsworth, V. Wright and D. Dowson, 1975. Quantitative study of stiffness in the
knee joint. Annals of Rheumatoid Diseases 34, 286-291.
Tardieu, C., P. Colbeau-Justin, M.D. Bret, A. Lespargot and G. Tardieu, 1981. Effects on
torque-angle curve of differences between therecorded tibia-calcaneal angle and the true
anatomical angle. European Journal of Applied Physiology 46, 41-45.
Vincken. M.H., C.C.A.M. Gielen and J.J. Denier van der Gon, 1983. Intrinsic and afferent
components in apparent muscle stiffness in man. Neuroscience 9, 529-534.
Viviani, P., J.F. Soechting and C.A. Terzuolo, 1976. Influence of mechanical properties on the
relation between EMG activity and torque. Journal of Physiology (Paris) 72, 45-5X.
Walmsley, B. and U. Proske, 1981. Comparison of stiffness of soleus and media1 gastrocnemius
muscles in cats, Journal of Neurophysiology 46, 250-259.
692 M.L. Latash, KM. Zatsiorsky /Joint stiffness
Walsh, E.G. and G.W. Wright, 1982. A hanging-hand tremorograph. Journal of Physiology 329.
I-2P.
Weber, E., 1847. Wagners Handworterbuch der Physiologie 3, 81.
Weiss, P.L., I.W. Hunter and R.E. Kearney, 1988. Human ankle joint stiffness over the full range
of muscle activation levels. Journal of Biomechanics 21, 539-544.
Werestchagin, N., 1931. Untersuchung iiber Musketonus und Ermiidung (Neue Methode zur
Bestimmung des Dehnungswiderstandes des Muskelsl. Pfliiger’s Archiv fiir die gesamte
Physiologie 227, 1X8-201.
Wilkie, D.R., 1950. The relation between the force and velocity in human muscle. Journal of
Physiology 110, 249-280.
Wilkie, D.R., lY56a. The mechanical properties of muscle. British Medical Bulletin 12, 1777182.
Wilkie, D.R., 1956b. Measurement of the series elastic component at various times during a
single muscle twitch. Journal of Physiology 134, 527-530.
Wilkie, D.R., 1979. ‘General discussion’. In: H. Sugi and G.H. Pollach ted.), Cross-bridge
mechanism in muscle contraction (p. 6341. Baltimore: University Park Press.
Woledge, R.C., N.A. Curtin and E. Homsher, 198.5. Energetic aspects of muscle contraction.
Monographs of the Physiological Society No. 41. London: Academic Press.
Woo, S.L.-Y., 1986. ‘Biomechanics of tendons and ligaments’. In: G.W. Schmid-Schonbein,
S.L.-Y. Woo SL-Y and B.W. Zweifach Frontiers in biomechanics (pp. 180-195). Berlin:
Springer-Verlag.
Yamada, H., 1970. Strength of biological materials (Ed. by F.G. Evans). Baltimore: Williams and
Wilkins.
Yoneda, T., K. Oishi and A. Ishida, 1983. Variation of amount of muscle discharges during
ballistic isometric voluntary contraction in man. Brain Research 275, 305-309.
Yoon, Y.S. and J.M. Mansour, 1982. The passive elastic moment at the hip. Journal of
Biomechanics 15, 905-910.
Zahalac, G.I. and S.I. Heyman, 1979. A quantitative evaluation of the frequency-response
characteristics of active human skeletal muscle in viva. Journal of Biomechanical Engineering
101, 28-37.
Zajac, F.E., 1989. Muscle and tendon: Properties, models, scaling, and application to biomechan-
its and motor control. CRC Critical Reviews in Biomedical Engineering 17, 359-411.
Zatsiorsky (Zaciorskij), V.M., 1977. Die kijrperlichen Eigenschaften des Sportlers (3rd ed.1.
Berlin: Bartels u. Wernitz.
Zatsiorsky, V.M., A.S. Aruin, L.M. Raisin and G.J. Panovko, 1981. ‘The determination of the
equivalent biomechanical characteristics of the ankle joint muscles by vibration tests’. In: G.
Bianchi, K.V. Frolov and A. Oledzki teds.), Man under vibration. Suffering and protection.
International CISM-IFToMM-Symposium. Udine, Italy, April 336, 1979. Proceedings, PWN
(pp. 166-175). Amsterdam: Elsevier.
Zatsiorsky, V.M. and A.S. Aruin, 1984. Biomechanical characteristics of the human ankle joint
muscles. European Journal of Applied Physiology 52, 400-406.